Identification of polymorphism and association analysis with reproductive traits in the porcine RNF4 gene

The ring finger protein 4 gene (RNF4), which might play a role in fetal germ cell development as well as in oocyte and granulosa cell maturation, was one of the potential candidate genes for reproductive traits. In the present work, we isolated the complete coding sequence of porcine RNF4 gene, identified a single nucleotide polymorphism (SNP: T/C) in intron5, and developed a PCR-SacII-RFLP genotyping assay. Association of this SNP with reproductive traits was assessed in three populations with diverse genetic backgrounds. One was Chinese Qingping sows. Another was consisted of crossbred sows derived from Landrace, Large White, Chinese Tongcheng and/or Chinese Meishan (Line DIV). The third is Large White × Meishan (LW × M) F₂ slaughtered population. Statistical analysis demonstrated that, in the first parity, the difference between RNF4 genotypes and reproductive traits of both Qingping and Line DIV sows was not significant. In the second and subsequent litters, CC animals in Qingping population had more piglets born (+1.74 piglets) and piglets born alive (+2.02 piglets) than sows with the TT genotype (P < 0.05). Line DIV sows inheriting the CC genotype had additional 0.69 piglets born compared to the TC animals (P < 0.05) in second and subsequent litters. No significant difference was observed between genotypes and reproductive tracts components in F₂ animals. In addition, we found RNF4 gene has a significant additive effect on both piglet born and piglet born alive in Qingping animals (P < 0.05). Results here suggested that the RNF4 SNP was significantly associated with litter size in two populations and could be useful in selection for increasing litter size in pigs. Further studies were needed to confirm these preliminary researches.     

Impact of the ESR gene on litter size and production traits in Czech Large White pigs

To evaluate the effect of the PvuII polymorphism of the oestrogen receptor gene on litter size and production traits in Czech Large White swine, data from 1250 sows and 3600 litters were analysed with two four-trait animal models. The traits in the first model were number of piglets born alive in a sow's first litter, number of piglets born alive in second and subsequent litters, lifetime daily gain and lean meat percentage. The second model included number of piglets born, number of piglets born alive, number of piglets weaned and litter weight at weaning from first and subsequent litters. The oestrogen receptor (ESR) locus significantly affected prolicacy in the first parity and averaged over all parities (P < 0.05), with allele A superior to allele B. In the first parity, AA sows produced approximately 0.5 more live piglets per litter than BB sows. Averaged over all parities, this difference was c. 0.25 piglets. Results for total number of piglets born and number of piglets weaned were similar to results for numbers born alive. No significant dominance effect was found for prolificacy traits. For litter weight at weaning, no significant additive effect was observed at the ESR locus, but a significant negative dominance effect (-1.5 kg) was estimated averaged across parities (litters of AB sows were similar to litters of BB sows for this trait). No pleiotropic effect of the ESR polymorphism on average daily gain or lean meat percentage was found.     

SLA-11 mutations are associated with litter size traits in Large White and Chinese DIV pigs

Litter size is an important economic traits in pigs. SLA-11 gene is a member of SLA (swine leukocyte antigen) complex. In our previous study, the SLA-11 gene was differentially expressed in PMSG-hCG stimulated preovulatory ovarian follicles of Chinese Taihu and Large White sows. Here, we identified two mutations (c.754-132 T?>?C and c.1421?+?38 T?>?C) in SLA-11 gene and analyzed the associations of two SNPs with litter size traits in Large White (n?=?263) and DIV (n?=?117) sows. The results showed that in Large White pigs, SLA-11 c.754-132?CC sows produced 0.74 and 0.87 more pigs per litter for TNB and NBA of all parities than did TT sows (p?<?.05); In DIV pigs, SLA-11 c.754-132?CC sows produced 1.17 more pigs per litter for TNB of all parities than did TC sows (p?<?.05). In Large White pigs, SLA-11 c.1421?+?38?CC sows produced 0.9 more pigs per litter for TNB of all parities than did TT sows (p?<?.05), while in DIV pigs SLA-11 c.1421?+?38?CC sows produced 0.84 and 0.7 less pigs per litter for TNB and NBA of all parities than did TT sows (p?<?.05). Our research indicated that SLA-11 mutations were potential molecular markers for improving the litter size traits in pigs.     

Effect of polymorphism in the leukemia inhibitory factor gene on litter size in Large White pigs

DNA polymorphism of the porcine leukemia inhibitory factory (LIF) was investigated and used to study the effects on litter size in Large White pigs. A total of 2,167 litter records from 420 sows genotyped at two SNP loci (LIF1 and LIF2) within LIF gene were analyzed to determine whether LIF influenced total number born (TNB) and number born alive (NBA). The results indicated that B allele at LIF1 locus and A allele at LIF2 locus seem to have advantageous effects on litter size. However, the combined analyzed results demonstrated that genotype AAAA, ABBB, and BBBB are better than genotype AAAB, AABB, and ABAB for TNB and NBA in either third to eighth parity or all parities. In all parities, the sows with AAAA genotype had an advantage of 1.76 piglets (P < 0.001) for TNB and 1.44 piglets (P < 0.01) for NBA per litter over the AAAB sows, respectively. The results in this study demonstrated that LIF gene was significantly associated with litter size in pigs. H. C. Lin and G. F. Liu contributed equally to this work.     

Effect of VNTR polymorphism of the Muc1 gene on litter size of pigs

An investigation was undertaken to study the association between the variable number of tandem repeats polymorphism of the Muc1 gene and the litter size in pigs. Four different alleles were found in three breeds. The sequence analysis shows that the repetitive region of pig Muc1 gene is an array of 108-bp repeats. A total of 2,430 litter records from 897 sows genotyped at Muc1 gene were used to analyze the total number born (TNB) and number born alive (NBA). The study of the effects on litter size suggests that TNB and NBA of genotype AA are the highest in Large White, and the TNB and NBA of the third to ninth parities are 1.61 and 2.29 piglets per litter higher (P < 0.05) than those of the genotype DD, respectively. In Landrace, TNB and NBA of the genotype AA are 1.68 (P < 0.01) and 1.58 (P < 0.05) piglets per litter higher than those of the BB genotype in the third to ninth parities, but for all parities the TNB of genotype AA were 0.76 piglets per litter (P < 0.05) higher than BB. In Duroc, the TNB and NBA of genotype AA are about 1.5 piglets per litter more than those of DD in the third to ninth parities, though not significantly. The research suggests that the smaller allele tends to have higher litter size. The results indicate that Muc1 gene is significantly associated with litter size in pigs.     

A SNP in the miR-27a gene is associated with litter size in pigs

MicroRNAs (miRNAs) are 20–25 nt, endogenous non-coding RNA molecules that act by binding to the complementary sequence of target messenger RNAs. Many evidences showed that miRNAs were involved in the process of germ proliferation and differentiation. In the present study, miR-27a gene was selected as a candidate gene for litter size due to its biological function, its location near a mummified pigs QTL, and its differentially expressed profile in Large White and Chinese Erhualian PMSG-hCG stimulated preovulatory ovaries. By comparative sequencing of miR-27a gene in Large White and Chinese Meishan pigs, one SNP (T/C) which created an additional HpaII site was detected. Then associations of this SNP with litter size traits were assessed in Large White (n = 142) and DIV (n = 140) pig populations. The statistical analysis demonstrated that AA differed from AB (P < 0.01) and BB (P < 0.05) for total number of piglets born in the first parities, and also differed from AB (P < 0.01) for the number of piglets born alive in all parities (P < 0.05) in DIV pigs. No significant difference was observed between different genotypes in Large White pigs.     

Association of four new single-nucleotide polymorphisms in follicle-stimulating hormone receptor and zona pellucida glycoprotein with reproductive traits in pigs

Two new single-nucleotide polymorphisms (SNPs) (C1166T and G1190A) were discovered in the follicle-stimulating hormone receptor (FSHR) gene and two (G261A and T302C) in the zona pellucida glycoprotein (ZP3) gene. These SNPs were genotyped in three Chinese domestic purebred sow lines (42 Small Meishan, 46 Qingping and 41 Jinhua sows) and three European purebred sow lines (225 Duroc, 195 Large White and 65 Landrace sows) by using SNP chips. Phenotypic data including the functional teat number (i.e. milk-producing teats, TN) and number of piglets born alive per litter (NBA). These traits were tested for association with the genotypes of four SNPs. The association analysis revealed genotype of G261A in the ZP3 gene was significantly (P < 0.01) associated with overall NBA and NBA at later parities (NBA2+) but not with NBA at first parity (NBA1). There was a significant (P < 0.05) difference between sows with genotype GG (14.83 ± 0.18) and AA (14.26 ± 0.09) in TN at position 261 in the ZP3 gene. No significant associations were observed for the SNPs in the FSHR gene with NBA or TN in our populations. The results showed that the new SNPs in the ZP3 gene may be an effective potential marker to be used in conjunction with traditional selection methods.     

Effect of polymorphism in the peroxisome proliferator-activated receptor gamma gene on litter size of pigs

The association of polymorphisms in peroxisome proliferator-activated receptor γ (PPARγ) gene with litter size was studied in Large White and Landrace pig. Three SNP loci (P1, P2 and P7) on PPARγ₂ gene were determined by PCR–SSCP and the results showed that there were A → G mutations at 220 and 324 bp in 5′-regulator region and at 147 bp in exon 6, respectively. Allele frequencies were analysed in two breeds. Information on 2341 litter records from 564 sows was used to analyse the trait total number born (TNB) and number born alive (NBA). In Large White, TNB and NBA of genotype BB for P2 locus were the lowest, and the TNB and NBA of third and following parities and all parities were 0.74 and 0.51 piglets per litter less (P < 0.001) than those of the highest genotype AB, respectively, but for P1 and P7 locus the beneficial genotype AA were more 0.4–0.8 piglets per litter (P < 0.05) than the inferior genotype AB. In landrace, TNB and NBA of the first parity of genotype BB for P1 locus were 2.0 piglets per litter higher than AA (P < 0.05), but for all parities the TNB and NBA of genotype BB were 0.66 and 0.97 piglets per litter (P < 0.05) higher than AA, respectively. At P2 locus, the TNB and NBA of the second parity of genotype AA were obviously higher than those of AB (P < 0.05). And at P7 locus, the TNB and NBA of each parity of genotype AA were both about 2 piglets per litter more than those of BB (P < 0.05). The results indicated that PPARγ gene was significantly associated with litter size in pigs.     

Effect of birth litter size, birth parity number, growth rate, backfat thickness and age at first mating of gilts on their reproductive performance as sows

The present study was performed to evaluate retrospectively the influence of birth litter size, birth parity number, performance test parameters (growth rate from birth to 100kg body weight and backfat thickness at 100kg body weight) and age at first mating (AFM) of gilts on their reproductive performance as sows. Traits analysed included remating rate in gilts (RRG), litter size, weaning-to-first-service interval (WSI), remating rate in sows and farrowing rate (FR). Data were collected from 11 Swedish Landrace (L) and 8 Swedish Yorkshire (Y) nucleus herds and included 20712 farrowing records from sow parities 1-5. Sows that farrowed for the first time during 1993-1997, having complete records of performance test and AFM, were followed up to investigate their subsequent reproductive performance until their last farrowing in 1999. Analysis of variance and multiple regression were applied to continuous data. Logistic regression was applied to categorical data. The analyses were based on the same animals and the records were split into six groups of females, i.e. gilts, primiparous sows, and sows in parities 2-5, respectively. Each additional piglet in the litter in which the gilt was born was associated with an increase of her own litter size of between 0.07 and 0.1 piglets per litter (P<0.001). Gilts born from sow parity 1 had a longer WSI as primiparous sows compared with gilts born from sow parity 4 (0.3 days; P<0.05) or parity 5 (0.4 days; P<0.01). Gilts with a higher growth rate of up to 100kg body weight had a larger litter size (all parities 1-5; P<0.05), shorter WSI (all parities 1-5; P<0.05) and higher FR (parities 2 and 5; P<0.05) than gilts with a lower growth rate. Gilts with a high backfat thickness at 100kg body weight had a shorter WSI as primiparous sows (P<0.001) compared with low backfat gilts, and 0.1 piglets per litter more as second parity sows (P<0.01). A 10 day increase in AFM resulted in an increase in litter size of about 0.1 piglet for primiparous sows (P<0.001) and a decrease (P<0.05) for sow parities 4 and 5.     

Variability and repeatability in gestation length related to litter performance in female pigs on commercial farms

Forecasting gestation length (GL) in sows enables producers to be prepared to provide assistance at farrowing, or for timely treatment to induce parturition. The objectives of the present study were to determine GL across parities, the repeatability and correlation of the GL, and associations of GL with the three litter size variables (total pigs born, pigs born alive, and dead piglets) and longevity. This study was conducted on 94 farms and encompassed 66,254 farrowing records of 13,715 sows born during 1999. Variance components and correlation analyses were used to determine the repeatability and the correlations of GL. Mixed-effects models were used to analyze associations of GL with litter size variables and longevity. The mean of GL across parities was from 115.2 to 115.4 d. The proportions of GL 114, 115, and 116 d for all farrowing events were 19.2, 30.8, and 22.2%, respectively. The GL between parities were correlated (0.40/=117 d (P<0.01). Sows with GL相似文献   

Influence of some sow characteristics on within-litter variation of piglet birth weight

Within-litter variation of piglet birth weight (BW0) is associated with an increased piglet mortality and a high variability in pig weight at weaning and weight or age at slaughter. Data collected in two experimental herds were used to quantify within-litter variability in BW0 and to assess the influence of factors mainly related to the sow. Within 24 h after birth, piglets born alive were individually weighed and stillborn piglets were collectively (first data set) or individually (second data set) weighed. The first data set was restricted to litters with no or only one stillborn piglet (3338 litters). It was used to assess the influence of genetic selection on BW0 variation by comparing litter characteristics before (1994 to 1996) and after (2001 to 2004) the development of hyperprolific sows in this herd. The second data set included all litters (n = 1596) from sows born between 2000 and 2004. For each litter, mean BW0 (mBW0) and its coefficient of variation (CVBW0) were calculated. Then, variance analyses were performed to test the influence of litter size, parity, year of sow birth and season at conception. Prolificacy improvement was associated with an increased CVBW0 in litters from pure Large White (LW) and Landrace × Large White (LR × LW) crossbred sows. The CVBW0 averaged 21% and was significantly influenced by litter size and parity. It increased from 15% to 24% when litter size varied from less than 10 piglets to more than 15 piglets. The proportion of small piglets (i.e. weighing less than 1 kg) increased concomitantly. The CVBW0 was not repeatable from a parity to the following. It was lowest for first and second parities (20%) and thereafter increased progressively. The CVBW0 was positively related to sow's backfat thickness gain during gestation. Taking into account litter size, parity, year of sow birth and season at conception explained 20% of BW0 variation. Thus, major part of heterogeneity is due to other factors, presumably including embryo genotype, on the one hand, and factors that influence embryo and foetus development, such as epigenetic factors, on the other hand.     

Molecular characterization and SNPs analysis of the porcine Deleted in AZoospermia Like (pDAZL) gene

The Deleted in AZoospermia Like (DAZL) gene is expressed in prenatal and postnatal germ cells. In this study, we cloned and characterized the porcine Deleted in AZoospermia Like (pDAZL) gene. We found the full-length coding sequence of the pDAZL encoded a protein of 295 amino acids with a RNA recognition motif (amino acids 41-111) and a DAZ repeat (amino acids 167-120). The deduced protein sequence of pDAZL is 92.5% and 91.5% similar to those of human and bovine, respectively. PCR-MspI-RFLP and PCR-TaqI-RFLP were established to detect an A/G mutation in intron 7 and a C/A mutation in intron 9, respectively. Associations of two SNPs with litter size traits were assessed in Large White (n=275) and DIV (n=128) pig populations, and the statistical analysis demonstrated that CC produced 0.716 more (P<0.05) piglets born alive than CD genotypes in Large White pigs at TaqI locus (C/A mutation in intron 9), and the dominance effect was 0.304 pig per litter (P<0.05). This result suggests that the pDAZL gene might be a good candidate gene of litter size trait and provides some marker information for marker-assisted selection (MAS).     

The effect of a SNP in ESR gene on the reproductive performance traits in Polish sows

The aim of the experiment was to detect polymorphism in the ESR gene to determine associations between the genotype and litter size in Polish Large White and Landrace sows. Reproductive traits investigated were: total number of piglets born (TNB), number of piglets born alive (NBA) and number of piglets weaned (TW). The polymorphism in ESR gene was detected using the PCR-RFLP method, with specific primers and the restriction enzyme AvaI. Two different alleles of ESR gene were identified: alleles A (0.71) and B (0.29). The relationship between the ESR genotypes and TBN, NBA and NW were analyzed. The analysis showed in first parity sows statistically significant (P < 0.01) differences between sows carrying different ESR genotypes. The analysis of ESR gene showed that sows with BB genotype had the largest litter size compared to AB and BB sows, but the difference was statistically not significant.     

雌激素受体基因和长白猪繁殖性能相关研究

通过检测612头长白母猪共2 239窝ESR基因的PvuⅡ酶切多态性,分析了不同ESR基因型和长白母猪繁殖性状的相关,以确定ESR基因在猪育种应用的可能性.头胎母猪、第2胎母猪和3胎以上母猪的资料分开统计.群体中B等位基因的频率很低,但似然比检验结果显示群体的基因频率处于哈代-温伯格连锁平衡状态.第1胎BB基因型母猪的总产仔数显著高于AA型母猪(12.05±0.82 vs 10.19±0.24)(P＜0.05),但BB基因型母猪的初生仔猪重显著低于AA型母猪(1.23±0.07vs 1.42±0.02)(P＜0.05).第3胎以上资料合并,BB基因型母猪的总产仔数显著多于AA和AB基因型母猪(11.98±0.63 vs10.90±0.48/10.92±0.51)(P＜0.05),产活仔数显著高于AA型母猪(10.31±0.58 vs 9.43±0.45)(P＜0.05);AB基因型母猪初生仔猪重显著低于AA型母猪(1.44±0.04 vs 1.48±0.04)(P＜0.05).所有资料合并,BB基因型母猪的总产仔数极显著高于AB型母猪(11.63±0.52 VS 10.63±0.42)(P＜0.01),显著高于AA型母猪(11.63±0.52 vs 10.70±0.40)(P＜0.05);BB基因型母猪的产活仔数显著高于AB和AA基因型母猪(10.15±0.50 vs 9.33±0.39/9.41±0.41)(P＜0.05).其余情况下各基因型母猪间繁殖性状间差异不显著(P＞0.05).总之,BB基因型母猪的总产仔数和产活仔数优于其他基因型母猪,但仔猪初生重较低.ESR基因可以作为遗传标记,用于本群长白猪产仔数的选择.     

Chromosomal abnormalities in hypoprolific boars

Four new chromosomal rearrangements are reported in the domestic pig: 3 reciprocal translocations, rcp(4;12)(p13;q13) in a crossbred boar, rcp(1;7)(q17;q26) in a Large White purebred boar, rcp(1;6)(q17;q35) in a purebred synthetic paternal line boar, and a pericentric inversion inv(2)(p13q11) in a crossbred boar. The 1/7 reciprocal translocation and the pericentric inversion were detected in animals that had sired small litters. The effect of the 1/7 translocation was accurately determined: -4.5 piglets born per litter, i.e. -36%. Both the 1/6 and 1/7 reciprocal translocations were of maternal origin. All the chromosomal rearrangements were highlighted using GTG and/or RBG banding techniques. Chromosome painting experiments were also carried out to confirm the proposed hypotheses for the three reciprocal translocations.     

Effect of equine chorionic gonadotropin on weaning-to-first service interval and litter size of female swine

We evaluated the effect of PMSG on the weaning-to-first service interval, total litter size and born alive litter size in swine. Four doses of PMSG (0, 500, 750 and 1,000 IU) were administered intramuscularly after weaning to sows at 3 different farms, grouped by parities (1, 2 and 3 or higher) and 2 distinct time periods. The associations among main effects and response variables were assessed by analysis of variance. Polynomial orthogonal terms were used to adjust the estimates of weaning-to-first service interval, total litter size and born alive litter size for the interaction effect of parity and PMSG treatment. The weaning-to-first service interval did not differ across periods and farms (P>0.05), although the interval was shorter (P<0.05) for Parity 3+ sows (4.97 d) than for Parity 1 sows (5.29 d), with no other differences in intervals observed across parities (P>0.05). Time period did not influence litter size (P>0.05), but there were differences in litter size across farms (P<0.05). Both litter size traits were lower for Parity 1 sows than for higher parity sows (P<0.05), but there were no differences in litter size between Parity 2 and 3+ sows (P>0.05). Litter size increased with PMSG dose in both Parities 1 and 2 (P<0.05), but not in Parity 3+ (P>0.05). A significant quadratic effect (P<0.05) of PMSG treatment in weaning-to-first service interval was observed for both Parity 1 and 2 sows, with the shortest intervals occurring with the 750 IU dose for Parity 1 sows. Administration of PMSG after weaning was associated with a shortened weaning-to-first service interval in Parity 1 sows and increased litter size in Parity 1 and 2 sows.     

Molecular cloning and polymorphism of the porcine H2AFZ gene

H2A histone family, member Z (H2A.Z) is required for early mammalian development. In the present study, the 932 bp of full-length cDNA encoding a 128 amino-acid protein and the sequences of intron 2 to 4 of the porcine H2A histone family, member Z (pH2AFZ) gene were obtained. By comparative sequencing of pH2AFZ gene in Large White and Meishan pigs, a 4 bp deletion/insertion in intron 2 was detected and a PCR-Bsu15I-RFLP was established to detect this variation. In DIV (4th Dam line of Chinese lean-type new lines) pigs, the first-parity females with AA genotype had fewer piglets born alive (-2.64 and -1.83 piglets per litter) than those with AB (P < 0.01) and BB (P < 0.05) genotype. The additive allelic and dominance effect were estimated to be 0.92 (P < 0.05) and -0.87 piglets per litter (P < 0.01) for number of piglets born alive, respectively. This result suggests that the pH2AFZ gene might be a good candidate gene of litter-size trait and provides some marker information for marker-assisted selection.     

Reproductive performance of purebred landrace and Yorkshire sows in Thailand with special reference to seasonal influence and parity number

The purpose of this study was to analyze reproductive performance in purebred Landrace and Yorkshire sows with special reference to seasonal influence and parity number, under tropical conditions where day length is almost constant throughout the year. Data from three purebred sow herds in Thailand during the period from 1993 to 1996 were analyzed. The two breeds were present in all three herds. The analysis comprised records of 3848 Landrace sow litters and 2033 Yorkshire sow litters. The statistical models included the fixed effects of month, year, parity, breed of the sow, herd, and two-way interactions of breed-parity, breed-herd, breed-month, breed-year, parity-month, month-herd, year-herd and month-year. The random effect of sow within breed was included in all models. Analysis of covariance was performed to analyze the effect of temperature, humidity and heat index on number of total born per litter (NTB), weaning to first service interval (WSI) and farrowing rate (FR). Landrace sows had significantly higher NTB (0.6 piglets), number of live born per litter (0.5 piglets), and average birth weight (0.13 kg) than Yorkshire sows (P<0.001). Farrowing rate was 3.9% higher in Landrace sows than in Yorkshire sows (P<0.01). However, Yorkshire sows had significantly shorter WSI (P<0.001) and significantly higher proportion of sows served within 7 days after weaning (P<0.01) than Landrace sows. No breed differences were found in number of stillborn per litter and weaning to conception interval. Parity had significant effect on all reproductive parameters analyzed. Number of total born and live born per litter was significantly lower for sows farrowing during the rainy season than in other seasons. Farrowing rate was low for sows mated during the hot and rainy season. Weaning to service interval and WSI7 were prolonged for sows weaned during the hot and rainy season. Reproductive performance was significantly unfavorably influenced by elevated temperature and heat index after mating (NTB and FR) or during lactation (WSI).     

Detection of quantitative trait loci for reproduction and production traits in Large White and French Landrace pig populations (Open Access publication)

A genome-wide scan was performed in Large White and French Landrace pig populations in order to identify QTL affecting reproduction and production traits. The experiment was based on a granddaughter design, including five Large White and three French Landrace half-sib families identified in the French porcine national database. A total of 239 animals (166 sons and 73 daughters of the eight male founders) distributed in eight families were genotyped for 144 microsatellite markers. The design included 51 262 animals recorded for production traits, and 53 205 litter size records were considered. Three production and three reproduction traits were analysed: average backfat thickness (US_M) and live weight (LWGT) at the end of the on-farm test, age of candidates adjusted at 100 kg live weight, total number of piglets born per litter, and numbers of stillborn (STILLp) and born alive (LIVp) piglets per litter. Ten QTL with medium to large effects were detected at a chromosome-wide significance level of 5% affecting traits US_M (on SSC2, SSC3 and SSC17), LWGT (on SSC4), STILLp (on SSC6, SSC11 and SSC14) and LIVp (on SSC7, SSC16 and SSC18). The number of heterozygous male founders varied from 1 to 3 depending on the QTL.